What is both necessary and sufficient to maintain avoidance responding in the shuttle box?

To learn what maintains the frequency of shuttle box avoidance responses, male rats from the Berkeley S₁ strain, after 200 trials of standard discriminative avoidance training, were given 100 additional trials under one of four different conditions. Responding at the maximum rate was maintained when animals performed under the training conditions or when responses continued to terminate the warning signal immediately, even though shock was never given for failing to respond. In contrast, avoidance responding was reduced markedly if, and only if, trials were given in which the signal ceased to terminate immediately (i.e. it shut off either well before or well after a response). This decrement occurred even though avoidance responses continued to avert shock. Thus, under the conditions of this experiment prompt signal offset was both necessary and sufficient to maintain the occurrence of well-established shuttle box avoidance responses.     

Preference and response suppression under different correlations between shock and a positive reinforcer in rats

A lever-press response by rats was reinforced by food in two successively presented types of trial signalled by different discriminative stimuli. When responding was punished by a shock in one type of trial, a groups for which the shock always preceded the reinforcer by .5 sec (positive correlation) showed less suppression in those trials than a group which received the shock and food separated in time (negative correlation). When, in a second experiment, rats were given a choice between food alone in one end of a shuttle box and either positively or negatively correlated shock and food in the other on a concurrent schedule, the group receiving the negatively correlated shock showed a greater preference for the food alone end. On the basis of a third experiment in which a tone was substituted for the shock in the choice situation, it was argued that the effect of correlation was not simply due to the stimulus properties of the shock. A final experiment demonstrated that when shock punishment is administered during extinction of the lever-press response, the rate of extinction is slower if the shock has been previously paired with the food reinforcer. Pairing a shock with food seems to attenuate the intrinsic aversiveness of the shock through Pavlovian counterconditioning.     

The generality of learned helplessness in the rat

Two experiments were simultaneously conducted in which two different groups of 40 rats each were exposed to one of two different stressors. In both experiments half the subjects were pretreated with shock, half with underwater exposure. For each pretreatment stressor, half the subjects were allowed to escape, the other half were not. The experiments differed in the test task used. Approximately 24 hr after pretreatment, one-half the subjects from each pretreatment group received 20 water-escape trials in an underwater maze, the other half received 20 shock-escape trials in a two-way shuttle box. The subjects in each of the inescapable pretreatment conditions were slower to escape in the subsequent shock-escape and water-escape tasks when compared with subjects in the corresponding escapable pretreatment condition. The “learned helplessness” effect appeared to be no smaller when aversive stimuli were changed between pretreatment and test than when they remained the same.     

Choice of a terminating over a non-terminating signal in free-operant avoidance

Rats chose between two signalled avoidance schedules. Under one schedule, responses in the presence of the signal terminated it and resulted in avoidance of shock; in the other, responses in the presence of the signal resulted in shock avoidance but signal termination was delayed for 10 sec. Pressing a second (changeover) lever produced change from one schedule to the other for 1 min. Once this 1-min period timed out, subjects could remain under the schedule in effect or could reinstate the other schedule for another 1-min period. All four subjects continuously changed over from the non-terminating to the terminating signal schedule. Changeover responding was not maintained when the termination contingency was removed. When changeover responding resulted in a change from a terminating to a non-terminating signal schedule, changeover responding did not occur.     

Conditioned memory modulation, freezing, and avoidance as measures of amygdala-mediated conditioned fear

Three conditioned aversive responses were used to infer the existence of an unobservable central state of "conditioned fear," and the roles of certain amygdala subregions in producing these responses were investigated. Rats received tone-shock pairings in one compartment of a shuttle box and no tones or shocks in the other, distinctive, compartment. They were then trained to find food in one arm of a Y-maze. After the final training trial they were exposed to different sets of stimuli in the shuttle box with no shock. Twenty-four hours later rats that had received immediate posttraining exposure to the conditioned stimuli (in the shock-paired compartment) made significantly more correct responses on the Y-maze than rats that had been exposed to the neutral stimuli (in the no-shock compartment) or rats that had received delayed posttraining exposure to the conditioned stimuli. This constitutes a demonstration of posttraining memory modulation by conditioned aversive stimuli. Freezing increased during posttraining exposure to the conditioned stimuli compared to the neutral stimuli. When subsequently allowed to move freely between the two compartments, the rats in all groups also showed significant conditioned avoidance of the compartment containing the conditioned stimuli. In a second experiment the effects of lesions confined to specific parts of the amygdala on the three conditioned responses (memory modulation, freezing, avoidance) were tested. Lesions of the central nucleus impaired all three conditioned responses; lesions of the medial nucleus impaired conditioned modulation and avoidance. These lesions had no effect on freezing during the training trials. Lesions of the lateral and basolateral nuclei attenuated freezing during both training and testing. The findings suggest that the central and medial nuclei of the amygdala may be important parts of neural circuits mediating conditioned responses that constitute conditioned aversive states, but that conditioned freezing may be mediated independently.     

Transfer of learning across reinforcers: Appetitive discrimination learning between stimuli previously associated with shock

In each of two experiments rats learned a food-rewarded discrete-trial discrimination in which the correct response in a two-lever Skinner box was signalled by an external stimulus. For one (experimental) group of subjects, one of the stimuli had previously been associated with electric shock while the other had been used to signal the absence of shock. A control group had previously experienced the stimuli uncorrelated with shock. In both experiments its was found that the control subjects learned the discrimination more rapidly than experimental subjects. The relevance of these findings to theories of the acquired distinctiveness of cues and to accounts of interaction between appetitive and aversive motivational systems is discussed.     

Some factors involved in the comparison of response systems: acquisition, extinction, and transfer of head-poke and lever-press Sidman avoidance

Head poking, a suggested natural escape reaction to shock for the rat, was compared to lever pressing in a Sidman avoidance study. Both responses could be emitted at any time, but only one was effective in a given session. Acquisition and extinction of the two responses were compared under both signalled and unsignalled avoidance. Then, a test for transfer was conducted in which acquisition conditions were re-instated, but the effectiveness of the responses was reversed. Three differences between responses were noted: (a) head poking was superior in reducing shock rates under signalled conditions; (b) head poking was more resistant to extinction, especially under signalled conditions; (c) under unsignalled conditions, animals were unable to learn to head poke if they had previously learned to lever press. Findings a and c were pursued in later experiments. Finding a depended on the location of the warning signal with respect to the response system. When the lever press required approach to the warning signal, the head poke was superior. But when the head poke required approach to the warning signal, the two responses were equally effective. Finding c depended on the absence of feedback for head poke during transfer. Two conclusions are offered: first, the two responses appear to obey the same laws when their topographical differences are taken into account. Second, response feedback appears to be more critical in transfer than in original acquisition.     

Nonselective motor-level changes associated with selective response inhibition: evidence from response force measurements

In the present study, we examined the effects of selective response inhibition on motor production using response force measures within a task that was based on that of Aron and Verbruggen (Psychological Science, 19, 1146–1153, 2008). In each trial, participants were signaled to respond bimanually with the two index fingers or the two middle fingers. After a short delay, a stop signal was sometimes presented, indicating that one of the two finger responses should be withheld. A given response was slowed when the response on the other hand was stopped, replicating a previously observed stopping interference effect. In addition, the given response was also made more forcefully when the response on the other hand was stopped, indicating that the requirement to stop one activated response has global motor-level consequences for other responses that are to be carried out normally.     

Response control with titration of punishment

Pigeons and rats were exposed to multiple schedules with different schedules of electric shock superimposed on identical schedules of food reinforcement during each of two components. During one component, (adjusting-intensity) the intensity of electric shock depended on responding. Each response increased the intensity while intensity decreased between responses. During the other component (constant-intensity) the intensity was fixed at the value at which it had been adjusted at the end of the immediately preceeding adjusting-intensity component. In one experiment, shock was continuous during both components. In another experiment, instead of continuous shock, a brief pulse was delivered immediately after each response. During the adjusting-intensity component of both experiments, pigeons and rats responded at a rate just sufficient to keep the shock constant (critical rate). During the constant-intensity component, responding depended on whether shock was delivered continuously or in pulses. When shock was continuous, response rate during the constant-intensity component was higher than the critical rate. When shock was pulsed, response rate during the constant-intensity component was equal to the critical rate.     

Methylphenidate and stimulus control of avoidance behavior

The introduction of a warning signal that preceded a scheduled shock modified the temporal distribution of free-operant avoidance responses. With response-shock and shock-shock intervals held constant, response rates increased only slightly when the response-signal interval was reduced. The result is consistent with Sidman's (1955) findings under different conditions, but at variance with Ulrich, Holz, and Azrin's (1964) findings under similar conditions. Methylphenidate in graded doses increased response rates, modifying frequency distributions of interresponse times. Drug treatment may have disrupted a "temporal discrimination" formed within the signal-shock interval. More simply, methylphenidate influenced response rates by increasing short response latencies after signal onset; this effect was more prominent than the drug's tendency to increase the frequency of pre-signal responses. When signal-onset preceded shock by 2 sec, individual differences in performance were marked; methylphenidate suppressed responding in one rat as a function of increasing dose levels to a greater degree than in a second animal, but both subjects received more shocks than under control conditions.     

Schedules using noxious stimuli. III. Responding maintained with response-produced electric shocks

Responding was maintained in two squirrel monkeys under several variations of a 10-min fixed-interval schedule of electric shock presentation. The monkeys were first trained under a 2-min variable-interval schedule of food presentation, and then under a concurrent schedule of food presentation and shock presentation. In one monkey, when shocks (12.6 ma) followed each response during the last minute of an 11-min cycle ending with a timeout period, responding was increased during the first 10 min and suppressed during the last minute of each cycle. When the shock schedule was eliminated, both the enhancement and suppression disappeared, and a steady rate of responding was maintained under the variable-interval schedule. When the food schedule was eliminated, the shock schedule maintained a characteristic fixed-interval pattern of responding during the first 10 min, but suppressed responding during the last minute of each cycle. The fixed-interval pattern of responding was maintained when the timeout period was eliminated and when only one shock could occur at the end of the cycle. In the second monkey, responding under the concurrent food and shock schedule was suppressed when responses produced shocks after 3-min. Under an 11-min cycle, responding continued to be maintained at increasing shock intensities. When the food schedule was eliminated, a fixed-interval pattern of responding was maintained under a 10-min schedule of shock presentation (12.6 ma). Whether response-produced electric shocks suppressed responding or maintained responding depended on the schedule of shock presentation.     

Category learning in the color-word contingency learning paradigm

In the typical color-word contingency learning paradigm, participants respond to the print color of words where each word is presented most often in one color. Learning is indicated by faster and more accurate responses when a word is presented in its usual color, relative to another color. To eliminate the possibility that this effect is driven exclusively by the familiarity of item-specific word-color pairings, we examine whether contingency learning effects can be observed also when colors are related to categories of words rather than to individual words. To this end, the reported experiments used three categories of words (animals, verbs, and professions) that were each predictive of one color. Importantly, each individual word was presented only once, thus eliminating individual color-word contingencies. Nevertheless, for the first time, a category-based contingency effect was observed, with faster and more accurate responses when a category item was presented in the color in which most of the other items of that category were presented. This finding helps to constrain episodic learning models and sets the stage for new research on category-based contingency learning.     

Reactance and the dynamics of disagreement: multiple paths from threatened freedom to resistance to persuasion

Many experiments show that threats to attitudinal freedom create reactance, but the underlying dynamics of reactance‐based disagreement have not received much attention. The present experiments identified two paths from threats to disagreement. In one path, threats to attitudinal freedom directly motivate disagreement; in the other, negative cognitive responses mediate the threat's effect on disagreement. Two experiments demonstrated the causes and consequences of each path from threat to persuasion. When a communicator threatened freedom at the beginning of the message, unfavorable cognitive responses (counterarguing, negative perceptions of the source's credibility) fully mediated the effect of threat on disagreement. When the threat appeared at the end of the message however, threat had a direct, unmediated effect on disagreement (Experiment 1). The two paths had different consequences for sleeper effects: disagreement rooted in negative cognitive responses persisted, whereas disagreement directly motivated by the threat declined when the threat was removed (Experiment 2). Implications for reactance and for threat‐based sleeper effects are discussed. Copyright © 2006 John Wiley & Sons, Ltd.     

Complex effects of NMDA receptor antagonist APV in the basolateral amygdala on acquisition of two-way avoidance reaction and long-term fear memory

Although much has been learned about the role of the amygdala in Pavlovian fear conditioning, relatively little is known about an involvement of this structure in more complex aversive learning, such as acquisition of an active avoidance reaction. In the present study, rats with a pretraining injection of the N-methyl-D-aspartate (NMDA) receptor antagonist, 2-amino-5-phosphonopentanoic acid (APV), into the basolateral amygdala (BLA) were found to be impaired in two-way active avoidance learning. During multitrial training in a shuttle box, the APV-injected rats were not different from the controls in sensitivity to shock or in acquisition of freezing to contextual cues. However, APV injection led to impaired retention of contextual fear when tested 48 h later, along with an attenuation of c-Fos expression in the amygdala. These results are consistent with the role of NMDA receptors of the BLA in long-term memory of fear, previously documented in Pavlovian conditioning paradigms. The APV-induced impairment in the active avoidance learning coincided with deficits in directionality of the escape reaction and in attention to conditioned stimuli. These data indicate that normal functioning of NMDA receptors in the basolateral amygdala is required during acquisition of adaptive instrumental responses in a shuttle box but is not necessary for acquisition of short-term contextual fear in this situation.     

Defensive burying, nest relocation, and pup transport in lactating female rats

A wire-wrapped wooden dowel was inserted through the wall of the nest compartment of a two-compartment box. There were four conditions. Some lactating female rats were shocked by the dowel when they first touched it, and some were not, after which the dowel was either immediately withdrawn from the chamber or left in place for the duration of the ensuing 30-min test period. Three defensive behaviours were observed during the tests: the mothers buried the shock source with bedding from the floor of the chamber, they transported their pups to the adjoining “safe” chamber, and they built a new nest in the safe chamber from material salvaged from their original nest. The shocked mothers that were confronted with the shock source throughout the test period displayed significantly more of each of these three defensive behaviours than did the mothers in the other three conditions. The methods used in this study provide a simple, reliable, safe paradigm for studying maternal defensive behaviour; the results indicate that the defensive capacities of the rat extend far beyond stereotypical flight, freeze, and fight responses, which have been the focus of most research on rodent defense.     

Some effects of methylphenidate on stimulus control of ratio avoidance behavior in the rat

After exposure to an avoidance schedule which included a warning signal, a rat was placed on a multiple schedule in which the first component was the same as before, i.e., a single response reset the response-shock interval, delaying shock, and the second component differed only in that four bar-presses were required to postpone shock. A fixed ratio requirement of four responses (FR 4) generated behavior resembling a fixed ratio requirement of one response (FR 1) since responding was controlled by the warning signal but more shocks were received. At a dosage of 2 mg/kg, methylphenidate given intraperitoneally decreased shock frequency during FR 4 periods while FR 1 behavior was not affected; at 4 mg/kg, stimulus control of avoidance responding was impaired during both components. Results at 4 mg/kg were partially confirmed by two animals exposed to an FR 4 avoidance schedule which included a warning signal but with different parameters. Response distributions showed that methylphenidate increased response rates in the absence of the warning signal, i.e., stimulus control of ratio-avoidance behavior was impaired although the increased response rates reduced shock frequency. One hour later responses again occurred more frequently during the signal than in its absence but shocks were less frequent than during control (non-drug) periods.     

Second-order schedules: brief shock at the completion of each component

Pigeons worked on second-order schedules in which completion of fixed-interval component schedules was reinforced with food according to a variable-interval schedule of reinforcement. The completion of each fixed-interval component resulted in the presentation of a brief electric shock. In one condition (shock-paired), the completion of every fixed-interval component, including those that ended in food, resulted in the shock. In another condition (shock-nonpaired), completion resulted in shock except for those components that ended in food. Shock presentations resulted in a positively accelerated rate within fixed-interval components. This patterning within components was similar whether the shock was intermittently paired with food or not. Response rates tended to decrease as shock intensity increased. The characteristic fixed-interval response pattern within components did not occur when shock presentations were omitted at the end of each component (tandem schedule). When shocks were scheduled but food was no longer presented (extinction) response rates declined to a near-zero level. The performance under shock conditions is similar to that in other studies in which visual and auditory stimuli are presented at the completion of component schedules.     

Duration-reduction of avoidance sessions as negative reinforcement

Five rats were exposed to a shock-postponement procedure in which responses on each of two levers initially had equivalent effects. After an initial training sequence that ensured at least some responding on each lever, an additional consequence was made conjointly operative on the previously less-preferred lever for each animal. Each response on this lever continued to postpone shock, but also reduced the session duration by one minute. The conjoint contingencies were operative until, through session-shortening responses and the passage of time, the session was scheduled to end in two minutes; during the final two minutes the session-shortening contingency was disabled while the shock-postponement contingency continued to be operative on both levers. When responding shifted to a predominance on the session-shortening lever, the conjoint contingency was shifted to the other lever; for four of the five rats this reversal was followed by two additional reversals. Two of the rats' responding showed clear, strong, and unambiguous sensitivity to the session-shortening contingency. The responding of two others was also systematically controlled by that contingency, but the effects were less clearcut. The fifth animal showed an initial shift when session-shortening was introduced, but its subsequent behavior proved insensitive to reversals of procedure. The results clearly indicate a sensitivity of behavior to events on a time scale quite distinct from that of immediate consequences. They also support an interpretation of avoidance sessions, considered in their entirety, as events whose contingent relationship to behavior can affect that behavior—even in the absence of stimuli that delineate those relationships. Finally, these results support an interpretation of aversively based conditioning within a broader context, analogous to the “open versus closed economy” interpretation of appetitively controlled behavior.     

Interaction of Pavlovian conditioning with a zero operant contingency: chronic exposure to signaled inescapable shock maintains learned helplessness effects

In four experiments we used triads, consisting of escapable-shock (ES), yoked inescapable-shock (IS), and no-shock (NS) rats, to investigate the effect of the interaction between Pavlovian contingencies and a zero operant contingency (i.e., uncontrollability) upon subsequent shock-escape acquisition in the shuttle box. After exposure to 50 signals and shocks per session for nine sessions, interference with shuttle box escape acquisition for IS rats was a monotonically increasing function of the percentage of signal-shock pairings during training (Experiment 1), with 50% pairings producing little or no impairment. Without regard to signaling, ES rats performed as well as NS rats. Experiment 2 demonstrated that our training and test conditions led to substantial and equal impairment in IS rats preexposed for one session to 100% or 50% signal-shock pairings or to unsignaled shocks. In Experiment 3, chronic exposure to 100% signaled inescapable shocks resulted in impairment only if the signal (light) was present during the shuttle box test. The continuous presence of the signal during the test contrasted with its discrete (5-s) presentation during training and suggested that an antagonistic physiological reaction rather than a specific competing motor response had been conditioned. Experiment 4 provided evidence for possible conditioned opioid mediation by demonstrating contemporaneous stress-induced analgesia and shock-escape impairment in IS rats chronically exposed to 100%, but not to 50%, signal-shock pairings, and the elimination of both analgesia and escape interference by the opiate antagonist naltrexone. Thus, chronic exposure to uncontrollable shocks appears to maintain the impairment produced by acute exposure only if the shocks are adequately signaled.     

Habituation of the hiding response to cat odor in rats (Rattus norvegicus)

Cat odor-induced hiding was examined in rats (Rattus norvegicus) using an apparatus with a "hide box" at one end and a piece of a worn cat collar at the other end. Rats spent most of their time hiding on exposure to the cat collar, but this response gradually habituated over repeated daily exposures. Hiding was reversed by administering the anxiolytic drug midazolam (0.375 mg/kg). Rats showed increased anxiety on the elevated plus-maze after exposure to the collar. This response was absent in habituated rats, suggesting that habituation of hiding reflects decreased odor-induced anxiety. It was established that rats located in the hide box of the apparatus might not have detected the odor of the cat collar placed at the other end of the apparatus. This implies that habituation occurs after relatively modest levels of odor exposure. Overall, the results cast some doubt on claims that predatory odors in rats are akin to phobic stimuli in humans.